At frequencies above 3 kHz, the tympanic membrane vibrates chaotically. By having many resonances, the eardrum can transmit the broadest possible bandwidth of sound with optimal sensitivity. In essence, the eardrum works best through discord. The eardrum's success as an instrument of hearing can be directly explained through a combination of its shape, angular placement, and composition. The eardrum has a conical asymmetrical shape, lies at a steep angle with respect to the ear canal, and has organized radial and circumferential collagen fiber layers that provide the scaffolding. Understanding the role of each feature in hearing transduction will help direct future surgical reconstructions, lead to improved microphone and loudspeaker designs, and provide a basis for understanding the different tympanic membrane structures across species. To analyze the significance of each anatomical feature, a computer simulation of the ear canal, eardrum, and ossicles was developed. It is shown that a cone-shaped eardrum can transfer more force to the ossicles than a flat eardrum, especially at high frequencies. The tilted eardrum within the ear canal allows it to have a larger area for the same canal size, which increases sound transmission to the cochlea. The asymmetric eardrum with collagen fibers achieves optimal transmission at high frequencies by creating a multitude of deliberately mistuned resonances. The resonances are summed at the malleus attachment to produce a smooth transfer of pressure across all frequencies. In each case, the peculiar properties of the eardrum are directly responsible for the optimal sensitivity of this discordant drum.collagen fibers ͉ hearing sensitivity ͉ middle ear ͉ transducers T he function of the middle ear in terrestrial mammals is to transfer acoustic energy between the air of the ear canal to the fluid of the inner ear. The first and crucial step of the transduction process takes place at the tympanic membrane, which converts sound pressure in the ear canal into vibrations of the middle ear bones. Understanding how the tympanic membrane manages this task so successfully over such a broad range of frequencies has been a subject of research since Helmholtz's publication in 1868 (1, 2).Even though the function of the eardrum is clear and the anatomy of the eardrum is well characterized, the connection between the anatomical features and the ability of the eardrum to transduce sound has been missing. The missing structurefunction relationships can be summarized by the following three questions. Why does the mammalian eardrum have its distinctive conical and toroidal shape? What is the advantage of its angular placement in the ear canal? What is the significance of its highly organized radial and circumferential fibers?The shape of the human and feline eardrum is known from detailed Moiré interferometry contour maps (refs. 3 and 4 and Fig. 1a). From the contour maps, three-dimensional reconstructions reveal the striking similarity of the two eardrums. In both cases, the eardrum has an ell...